Christophe Boesch

Cultures in chimpanzees

As an increasing number of field studies of chimpanzees (Pan troglodytes) have achieved long-term status across Africa, differences in the behavioural repertoires described have become apparent that suggest there is significant cultural variation. Here we present a systematic synthesis of this information from the seven most long-term studies, which together have accumulated 151 years of chimpanzee observation. This comprehensive analysis reveals patterns of variation that are far more extensive than have previously been documented for any animal species except humans. We find that 39 different behaviour patterns, including tool usage, grooming and courtship behaviours, are customary or habitual in some communities but are absent in others where ecological explanations have been discounted. Among mammalian and avian species, cultural variation has previously been identified only for single behaviour patterns, such as the local dialects of song-birds,. The extensive, multiple variations now documented for chimpanzees are thus without parallel. Moreover, the combined repertoire of these behaviour patterns in each chimpanzee community is itself highly distinctive, a phenomenon characteristic of human cultures but previously unrecognised in non-human species.

Fission-Fusion Dynamics: New Research Frameworks

Renewed interest in fission‐fusion dynamics is due to the recognition that such dynamics may create unique challenges for social interaction and distinctive selective pressures acting on underlying communicative and cognitive abilities. New frameworks for integrating current knowledge on fission‐fusion dynamics emerge from a fundamental rethinking of the term “fission‐fusion” away from its current general use as a label for a particular modal type of social system (i.e., “fission‐fusion societies”). Specifically, because the degree of spatial and temporal cohesion of group members varies both within and across taxa, any social system can be described in terms of the extent to which it expresses fission‐fusion dynamics. This perspective has implications for socioecology, communication, cognitive demands, and human social evolution.

Tool Use and Tool Making in Wild Chimpanzees

Reported incidences of tool use and tool making for three wild chimpanzee populations increase from Mahale (12 and 3 types of use and making, respectively), Gombe (16 and 3) to Taï (19 and 6). Sticks are commonly used and prepared at all three sites. However, Taï chimpanzees seem to perform more modifications on the material before using it. They are also the only chimpanzees seen to pound objects with tools and to combine two different tool uses to get access to one food item. Tool making is the rule for abundant material (grass, twigs), but appears to be rarer for scarce, hard material (clubs, stones). Factors involved in the acquisition and the benefit of tool use are discussed along with factors affecting the frequency and complexity of tool making in chimpanzees.

Quantitative polymerase chain reaction analysis of DNA from noninvasive samples for accurate microsatellite genotyping of wild chimpanzees (Pan troglodytes verus)

Noninvasive samples are useful for molecular genetic analyses of wild animal populations. However, the low DNA content of such samples makes DNA amplification difficult, and there is the potential for erroneous results when one of two alleles at heterozygous microsatellite loci fails to be amplified. In this study we describe an assay designed to measure the amount of amplifiable nuclear DNA in low DNA concentration extracts from noninvasive samples. We describe the range of DNA amounts obtained from chimpanzee faeces and shed hair samples and formulate a new efficient approach for accurate microsatellite genotyping. Prescreening of extracts for DNA quantity is recommended for sorting of samples for likely success and reliability. Repetition of results remains extensive for analysis of microsatellite amplifications beginning from low starting amounts of DNA, but is reduced for those with higher DNA content.

Microsatellite scoring errors associated with noninvasive genotyping based on nuclear DNA amplified from shed hair

In the context of a study of wild chimpanzees, Pan troglodytes verus, we found that genotypes based on single PCR amplifications of microsatellite loci from single shed hair have a high error rate. We quantified error rates using the comparable results of 791 single shed hair PCR amplifications of 11 microsatellite loci of 18 known individuals. The most frequent error was the amplification of only one of the two alleles present at a heterozygous locus. This phenomenon, called allelic dropout, produced false homozygotes in 31% of single‐hair amplifications. There was no difference in the probability of preferential amplification between longer and shorter alleles. The probability of scoring false homozygotes can be reduced to below 0.05 by three separate amplifications from single hairs of the same individual or by pooling hair samples from the same individual. In this study an additional 5.6% of the amplifications gave wrong genotypes because of contamination, labelling and loading errors, and possibly amplification artefacts. In contrast, amplifications from plucked hair taken from four dead individuals gave consistent results (error rate < 0.01%, n= 120). Allelic dropout becomes a problem when the DNA concentration falls below 0.05 ng/10 μL in the template as it can with shed hair, and extracts from faeces and masticated plant matter.

CHARTING CULTURAL VARIATION IN CHIMPANZEES

Cultural variation among chimpanzee communities or unit-groups at nine long-term study sites was charted through a systematic, collaborative procedure in which the directors of the sites first agreed a candidate list of 65 behaviour patterns (here fully defined), then classified each pattern in relation to its local frequency of occurrence. Thirty-nine of the candidate behaviour patterns were discriminated as cultural variants, sufficiently frequent at one or more sites to be consistent with social transmission, yet absent at one or more others where environmental explanations were rejected. Each community exhibited a unique and substantial profile of such variants, far exceeding cultural variation reported before for any other non-human species. Evaluation of these pan-African distributions against three models for the diffusion of traditions identified multiple cases consistent with cultural evolution involving differentiation in form, function and targets of behaviour patterns.

Mental map in wild chimpanzees: An analysis of hammer transports for nut cracking

The mental map of wild chimpanzees is analyzed in the context of their transports of clubs and stones used for cracking two species of nuts of different hardness,Coula edulis andPanda oleosa, in the Tai National Park (Ivory Coast). For the harderPanda nuts, they transport the harder hammers, i.e., almost exclusively stones, hammers of greater weight, and the transports are longer than forCoula nuts. The analysis made for the transports forPanda nuts shows that they seem to remember the location of stones and to choose the stones so as to keep the transport distance minimal. The chimpanzees seem to possess an Euclidian space, which allows them to somehow measure and remember distances; to compare several such distances so as to choose the stone with the shortest distance to a goal tree; to correctly locate a new stone location with reference to different trees; and to change their reference point so as to measure the distance to eachPanda tree from any stone location. They also combine the weight and the distance. The wild chimpanzees of the Tai National Park seem to possess concrete operation abilities in spatial representation.

Factors affecting the amount of genomic DNA extracted from ape faeces and the identification of an improved sample storage method

Genetic analysis using noninvasively collected samples such as faeces continues to pose a formidable challenge because of unpredictable variation in the extent to which usable DNA is obtained. We investigated the influence of multiple variables on the quantity of DNA extracted from faecal samples from wild mountain gorillas and chimpanzees. There was a small negative correlation between temperature at time of collection and the amount of DNA obtained. Storage of samples either in RNAlater solution or dried using silica gel beads produced similar results, but significantly higher amounts of DNA were obtained using a novel protocol that combines a short period of storage in ethanol with subsequent desiccation using silica.

Optimisation of Nut-Cracking With Natural Hammers By Wild Chimpanzees

The chimpanzees of the Tai National Park, Ivory Coast, use sticks and stones to open 5 different species of nuts. In spite of an unfavourable availability of the material in the forest, the animals choose their tools adaptively. For cracking harder nuts, they use harder and heavier tools and transport tools more often and from farther away. Some aspects of the evolution of tool-use in primates are discussed.

Generation times in wild chimpanzees and gorillas suggest earlier divergence times in great ape and human evolution

Fossils and molecular data are two independent sources of information that should in principle provide consistent inferences of when evolutionary lineages diverged. Here we use an alternative approach to genetic inference of species split times in recent human and ape evolution that is independent of the fossil record. We first use genetic parentage information on a large number of wild chimpanzees and mountain gorillas to directly infer their average generation times. We then compare these generation time estimates with those of humans and apply recent estimates of the human mutation rate per generation to derive estimates of split times of great apes and humans that are independent of fossil calibration. We date the human–chimpanzee split to at least 7–8 million years and the population split between Neanderthals and modern humans to 400,000–800,000 y ago. This suggests that molecular divergence dates may not be in conflict with the attribution of 6- to 7-million-y-old fossils to the human lineage and 400,000-y-old fossils to the Neanderthal lineage.